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Microbats constitute the
suborder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and recognized ...
Microchiroptera within the order Chiroptera (
bat Bats are flying mammals of the order Chiroptera (). With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most birds, flying with their very long spread-out ...
s). Bats have long been differentiated into Megachiroptera (megabats) and Microchiroptera, based on their size, the use of echolocation by the Microchiroptera and other features; molecular evidence suggests a somewhat different subdivision, as the microbats have been shown to be a
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
group.


Characteristics

Microbats are long. Most microbats feed on insects, but some of the larger species hunt birds, lizards, frogs, smaller bats or even
fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
. Only three species of microbat feed on the blood of large mammals or birds ("
vampire bat Vampire bats, members of the subfamily Desmodontinae, are Phyllostomidae, leaf-nosed bats currently found in Central and South America. Their food source is the blood of other animals, a dietary trait called hematophagy. Three extant bat species ...
s"); these bats live in South and Central America. Although most "Leaf-nose" microbats are fruit and nectar-eating, the name “leaf-nosed” isn't a designation meant to indicate the preferred diet among said variety. Three species follow the bloom of columnar cacti in northwest Mexico and the Southwest United States northward in the northern spring and then the blooming agaves southward in the northern fall (autumn). Other leaf-nosed bats, such as '' Vampyrum spectrum'' of South America, hunt a variety of prey such as lizards and birds. The horseshoe bats of Europe, as well as California leaf-nosed bats, have a very intricate leaf-nose for echolocation, and feed primarily on insects.


Differences from megabats

* Microbats use echolocation, whereas megabats do not typically. (The Egyptian fruit bat '' Rousettus egyptiacus'' is an exception, but does not use the larynx echolocation method of microbats, instead giving scientists the theory that it clicks using its nasal passages and back of its tongue.) * Microbats lack the claw at the second finger of the forelimb. This finger appears thinner and almost bonded by tissue with the third finger for extra support during flight. * Megabats lack tails, with the exception of a few genera such as ''Nyctimene'', whereas this trait only occurs in certain species of microbats. * The ears of microbats possess a tragus (thought to be crucial in echolocation) and are relatively larger than megabat ears, whereas megabat ears are comparatively small and lack a tragus. * Megabat eyes are quite large, whereas microbat eyes are comparatively smaller.


Dentition

The form and function of microbat teeth differ as a result of the various diets these bats can have. Teeth are primarily designed to break down food; therefore, the shape of the teeth correlate to specific feeding behaviors. In comparison to megabats which feed only on fruit and nectar, microbats illustrate a range of diets and have been classified as
insectivore file:Common brown robberfly with prey.jpg, A Asilidae, robber fly eating a hoverfly An insectivore is a carnivore, carnivorous animal or plant which eats insects. An alternative term is entomophage, which can also refer to the Entomophagy ...
s,
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they ar ...
s, sanguinivores,
frugivore A frugivore ( ) is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance ...
s, and
nectarivore In zoology, a nectarivore is an animal which derives its energy and nutrient requirements from a diet consisting mainly or exclusively of the sugar-rich nectar produced by flowering plants. Nectar as a food source presents a number of benefits ...
s. Differences seen between the size and function of the canines and molars among microbats in these groups vary as a result of this. The diverse diets of microbats reflect having dentition, or cheek teeth, that display a morphology derived from dilambdodont teeth, which are characterized by a W-shaped ectoloph, or stylar shelf. A W-shaped dilambdodont upper molar includes a metacone and paracone, which are located at the bottom of the “W”; while the rest of the “W” is formed by crests that run from the metacone and paracone to the cusps of the stylar self. Microbats display differences between the size and shape of their canines and molars, in addition to having distinctive variations among their skull features that contribute to their ability to feed effectively. Frugivorous microbats have small stylar shelf areas, short molariform rows, and wide palates and faces. In addition to having wide faces, frugivorous microbats have short skulls, which place the teeth closer to the fulcrum of the jaw lever, allowing an increase in jaw strength. Frugivorous microbats also possess a different pattern on their molars compared to carnivorous, insectivorous, nectarivorous, and sanguinivorous microbats. In contrast, insectivorous microbats are characterized by having larger, but fewer teeth, long canines, and shortened third upper molars; while carnivorous microbats have large upper molars. Generally, microbats that are insectivores, carnivores, and frugivores have large teeth and small palates; however, the opposite is true for microbats that are nectarivores. Though differences exist between the palate and teeth sizes of microbats, the proportion of the sizes of these two structures are maintained among microbats of various sizes.


Echolocation

Echolocation is the process where an animal produces a sound of certain wavelength, and then listens to and compares the reflected echoes to the original sound emitted. Bats use echolocation to form images of their surrounding environment and the organisms that inhabit it by eliciting ultrasonic waves via their
larynx The larynx (), commonly called the voice box, is an organ (anatomy), organ in the top of the neck involved in breathing, producing sound and protecting the trachea against food aspiration. The opening of larynx into pharynx known as the laryngeal ...
. The difference between the ultrasonic waves produced by the bat and what the bat hears provides the bat with information about its environment. Echolocation aids the bat in not only detecting prey, but also in orientation during flight.


Production of ultrasonic waves

Most microbats generate ultrasound with their larynx and emit the sound through their nose or mouth. Sound productions are generated from the
vocal folds In humans, the vocal cords, also known as vocal folds, are folds of throat tissues that are key in creating sounds through Speech, vocalization. The length of the vocal cords affects the pitch of voice, similar to a violin string. Open when brea ...
in mammals due to the elastic membranes that compose these folds. Vocalization requires these elastic membranes because they act as a source to transform airflow into acoustic pressure waves. Energy is supplied to the elastic membranes from the lungs, and results in the production of sound. The larynx houses the
vocal cords In humans, the vocal cords, also known as vocal folds, are folds of throat tissues that are key in creating sounds through Speech, vocalization. The length of the vocal cords affects the pitch of voice, similar to a violin string. Open when brea ...
and forms the passageway for the expiratory air that will produce sound. Microbat range in frequency from 14,000 to over 100,000
hertz The hertz (symbol: Hz) is the unit of frequency in the International System of Units (SI), often described as being equivalent to one event (or Cycle per second, cycle) per second. The hertz is an SI derived unit whose formal expression in ter ...
, well beyond the range of the human ear (typical human hearing range is considered to be from 20 to 20,000 Hz). The emitted vocalizations form a broad beam of sound used to probe the environment, as well as communicate with other bats. At the molecular level, it has been found that CPLX1 is involved in this ultrasonic wave production.


Laryngeally echolocating microbats

Laryngeal echolocation is the dominant form of echolocation in microbats, however, it is not the only way in which microbats can produce ultrasonic waves. Excluding non-echolocating and laryngeally echolocating microbats, other species of microbats and megabats have been shown to produce ultrasonic waves by clapping their wings, clicking their tongues, or using their nose. Laryngeally echolocating bats, in general, produce ultrasonic waves with their larynx that is specialized to produce sounds of short
wavelength In physics and mathematics, wavelength or spatial period of a wave or periodic function is the distance over which the wave's shape repeats. In other words, it is the distance between consecutive corresponding points of the same ''phase (waves ...
. The
larynx The larynx (), commonly called the voice box, is an organ (anatomy), organ in the top of the neck involved in breathing, producing sound and protecting the trachea against food aspiration. The opening of larynx into pharynx known as the laryngeal ...
is located at the
cranial Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Thi ...
end of the
trachea The trachea (: tracheae or tracheas), also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all animals' lungs. The trachea extends from ...
and is surrounded by cricothyroid muscles and thyroid cartilage. For reference, in
human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s, this is the area where the
Adam's apple The Adam's apple is the protrusion in the neck formed by the angle of the thyroid cartilage surrounding the larynx, typically visible in men, less frequently in women. The prominence of the Adam's apple increases in some men as a secondary mal ...
is located. Phonation of ultrasonic waves is produced through the vibrations of the vocal membranes in the expiratory air. The intensity that these vocal folds vibrate at varies with activity and between bat species. A characteristic of laryngeally echolocating microbats that distinguishes them from other echolocating microbats is the articulation of their stylohyal bone with their
tympanic bone The tympanic part of the temporal bone is a curved plate of bone lying below the squamous part of the temporal bone, in front of the mastoid process, and surrounding the external part of the ear canal. It originates as a separate bone (tympanic ...
. The stylohyal bones are part of the hyoid apparatus that help support the throat and larynx. The tympanic bone forms the floor of the
middle ear The middle ear is the portion of the ear medial to the eardrum, and distal to the oval window of the cochlea (of the inner ear). The mammalian middle ear contains three ossicles (malleus, incus, and stapes), which transfer the vibrations ...
. In addition to the connection between the stylohyal bone and the tympanic bone as being an indicator of laryngeally echolocating microbats, another definitive marker is the presence of a flattened and expanded stylohyal bone at the cranial end. Microbats that laryngeally echolocate must be able to distinguish between the differences of the pulse that they produce and the returning echo that follows by being able to process and understand the ultrasonic waves at a
neuron A neuron (American English), neurone (British English), or nerve cell, is an membrane potential#Cell excitability, excitable cell (biology), cell that fires electric signals called action potentials across a neural network (biology), neural net ...
al level, in order to accurately obtain information about their surrounding environment and orientation in it. The connection between the stylohyal bone and the tympanic bone enables the bat to neurally register the outgoing and incoming ultrasonic waves produced by the
larynx The larynx (), commonly called the voice box, is an organ (anatomy), organ in the top of the neck involved in breathing, producing sound and protecting the trachea against food aspiration. The opening of larynx into pharynx known as the laryngeal ...
. Furthermore, the stylohyal bones connect the larynx to the tympanic bones via a
cartilaginous Cartilage is a resilient and smooth type of connective tissue. Semi-transparent and non-porous, it is usually covered by a tough and fibrous membrane called perichondrium. In tetrapods, it covers and protects the ends of long bones at the joints ...
or fibrous connection (depending on the species of bat). Mechanically the importance of this connection is that it supports the larynx by anchoring it to the surrounding cricothyroid muscles, as well as draws it closer to the
nasal cavity The nasal cavity is a large, air-filled space above and behind the nose in the middle of the face. The nasal septum divides the cavity into two cavities, also known as fossae. Each cavity is the continuation of one of the two nostrils. The nas ...
during
phonation The term phonation has slightly different meanings depending on the subfield of phonetics. Among some phoneticians, ''phonation'' is the process by which the vocal folds produce certain sounds through quasi-periodic vibration. This is the defi ...
. The stylohyal bones are often reduced in many other mammals, however, they are more prominent in laryngeally echolocating bats and are part of the mammalian hyoid apparatus. The hyoid apparatus functions in breathing, swallowing, and phonation in microbats as well as other mammals. An important feature of the bony connection in laryngeally echolocating microbats is the extended articulation of the ventral portion of the tympanic bones and the proximal end of the stylohyal bone that bends around it to make this connection.


Classification

While bats have been traditionally divided into megabats and microbats, recent molecular evidence has shown the superfamily Rhinolophoidea to be more genetically related to megabats than to microbats, indicating the microbats are paraphyletic. To resolve the
paraphyly Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
of microbats, the Chiroptera were redivided into suborders Yangochiroptera (which includes Nycteridae, vespertilionoids, noctilionoids, and emballonuroids) and
Yinpterochiroptera The Yinpterochiroptera (or Pteropodiformes) is a suborder of the Chiroptera, which includes taxa formerly known as megabats and five of the microbat families: Rhinopomatidae, Rhinolophidae, Hipposideridae, Craseonycteridae, and Megadermatidae. T ...
, which includes megabats, rhinopomatids, Rhinolophidae, and Megadermatidae. This is the classification according to Simmons and Geisler (1998): Superfamily
Emballonuroidea Emballonuroidea is a Taxonomic rank, superfamily of bats containing two families: Emballonuridae and Nycteridae. It is one of three superfamilies in the suborder Yangochiroptera, the others being Noctilionoidea and Vespertilionoidea. Emballonuri ...
* Family Emballonuridae (
sac-winged bat Emballonuridae is a family (biology), family of microbats, many of which are referred to as sac-winged or sheath-tailed bats. They are widely distributed in tropical and subtropical regions around the world. The earliest fossil records are from ...
s or sheath-tailed bats) Superfamily Rhinopomatoidea * Family Rhinopomatidae ( mouse-tailed bats) * Family Craseonycteridae (bumblebee bat or
Kitti's hog-nosed bat Kitti's hog-nosed bat (''Craseonycteris thonglongyai''), also known as the bumblebee bat, is a near-threatened species of bat and the only extant member of the family Craseonycteridae. It occurs in western Thailand and southeast Myanmar, where it ...
) Superfamily
Rhinolophoidea Rhinolophoidea is a superfamily (taxonomy), superfamily of bats. It contains the following families: Craseonycteridae, Hipposideridae, Megadermatidae, Rhinolophidae, Rhinonycteridae, and Rhinopomatidae. It is one of two superfamilies that compris ...
* Family Rhinolophidae ( horseshoe bats) * Family Nycteridae ( hollow-faced bats or slit-faced bats) * Family Megadermatidae ( false vampires) Superfamily
Vespertilionoidea Vespertilionoidea is a superfamily of bats containing five families: Cistugidae, Miniopteridae, Molossidae, Natalidae, and Vespertilionidae. It is one of three superfamilies in the suborder Yangochiroptera, the others being Noctilionoidea ...
* Family Vespertilionidae (
vesper bat Vespertilionidae is a family of microbats, of the order Chiroptera, flying, insect-eating mammals variously described as the common, vesper, or simple nosed bats. The vespertilionid family is the most diverse and widely distributed of bat famili ...
s or evening bats) Superfamily Molossoidea * Family Molossidae (
free-tailed bat The Molossidae, or free-tailed bats, are a family of bats within the order Chiroptera. The Molossidae is the fourth-largest family of bats, containing about 110 species as of 2012. They are generally quite robust, and consist of many strong-fly ...
s) * Family Antrozoidae ( pallid bats) Superfamily Nataloidea * Family Natalidae ( funnel-eared bats) * Family
Myzopodidae ''Myzopoda'' is a genus of bat, the only member of the monotypic family (biology), family Myzopodidae, with two described species. Myzopodidae is unique as the only family of bats currently endemic to Madagascar. However, fossil discoveries indic ...
( sucker-footed bats) * Family
Thyropteridae Disk-winged bats are a small group of bats of the family Thyropteridae and genus ''Thyroptera''. They are found in Central and South America, usually in moist tropical rain forests. It is a very small family, consisting of a single genus with f ...
( disk-winged bats) * Family
Furipteridae Furipteridae is family of bats, allying two genera of single species, '' Amorphochilus schnablii'' (smoky bat) and the type '' Furipterus horrens'' (thumbless bat). They are found in Central and South America and are closely related to the ba ...
( smoky bats) Superfamily
Noctilionoidea Noctilionoidea is a Taxonomic rank, superfamily of bats containing seven families: Thyropteridae, Furipteridae, Noctilionidae, Mormoopidae, Phyllostomidae, Myzopodidae, and Mystacinidae. It is one of three superfamilies in the suborder Yangochir ...
* Family Noctilionidae ( bulldog bats or fisherman bats) * Family
Mystacinidae __NOTOC__ Mystacinidae is a family of unusual bats, the New Zealand short-tailed bats. There is one living genus, '' Mystacina'', with two species, one of which could have possibly become extinct in the 1960s. They are medium-sized bats, about ...
( New Zealand short-tailed bats) * Family
Mormoopidae The family Mormoopidae contains bats known generally as mustached bats, ghost-faced bats, and naked-backed bats. They are found in the Americas from the Southwestern United States to Southeastern Brazil. They are distinguished by the presen ...
(
ghost-faced bat The ghost-faced bat (''Mormoops megalophylla'') is a bat in the genus ''Mormoops''. It is one of only two extant species within its genus, the other being the much smaller ''Mormoops blainvillii''. They are nocturnal and hunt using Animal echolo ...
s or moustached bats) * Family Phyllostomidae ( leaf-nosed bats)


References


External links


Bat World Sanctuary


(''see "Recent publications"'')
Bat Conservation International
* {{Authority control Animals that use echolocation Bat taxonomy Paraphyletic groups